From around 4,000 to 2,000 BC the forest-steppe north-western Pontic region was occupied by people who shared a nomadic lifestyle, pastoral economy and barrow burial rituals. It has been shown that these groups, especially those associated with the Yamnaya culture, played an important role in shaping the gene pool of Bronze Age Europeans, which extends into present-day patterns of genetic variation in Europe. Although the genetic impact of these migrations from the forest-steppe Pontic region into central Europe have previously been addressed in several studies, the contribution of mitochondrial lineages to the people associated with the Corded Ware culture in the eastern part of the North European Plain remains contentious. In this study, we present mitochondrial genomes from 23 Late Eneolithic and Bronze Age individuals, including representatives of the north-western Pontic region and the Corded Ware culture from the eastern part of the North European Plain. We identified, for the first time in ancient populations, the rare mitochondrial haplogroup X4 in two Bronze Age Catacomb culture-associated individuals. Genetic similarity analyses show close maternal genetic affinities between populations associated with both eastern and Baltic Corded Ware culture, and the Yamnaya horizon, in contrast to larger genetic differentiation between populations associated with western Corded Ware culture and the Yamnaya horizon. This indicates that females with steppe ancestry contributed to the formation of populations associated with the eastern Corded Ware culture while more local people, likely of Neolithic farmer ancestry, contributed to the formation of populations associated with western Corded Ware culture.

Archaeological Centre Olomouc U Hradiska 42 6 779 00 Olomouc Czech Republic Department of History Section of Archaeology Philosophical faculty Palacký University Olomouc Na Hradě 5 771 80 Olomouc Czech Republic

Department of Human Evolutionary Biology Institute of Anthropology Faculty of Biology Adam Mickiewicz University in Poznan Umultowska 89 61 614 Poznań Poland

Department of Human Evolutionary Biology Institute of Anthropology Faculty of Biology Adam Mickiewicz University in Poznan Umultowska 89 61 614 Poznań Poland Laboratory of Genomics and Bioinformatics Institute of Molecular Genetics of the ASCR v v i Vídeňská 1083 142 20 Prague 4 Czech Republic

Human Evolution Department of Organismal Biology and SciLifeLab Uppsala University Norbyvägen 18C SE 752 36 Uppsala Sweden Centre for Anthropological Research University of Johannesburg Auckland Park 2006 Johannesburg South Africa

Institute of Archaeological Heritage Brno v v i Kaloudova 30 614 00 Brno Czech Republic

Institute of Archaeology Faculty of History Adam Mickiewicz University in Poznan Umultowska 89D 61 614 Poznań Poland

Institute of Archaeology Jagiellonian University Gołębia 11 31 007 Kraków Poland

Institute of Archaeology Maria Curie Skłodowska University Maria Curie Skłodowska Square 4 20 031 Lublin Poland

Laboratory of Morphology and Forensic Anthropology Department of Anthropology Faculty of Science Masaryk University Kotlářská 267 2 611 37 Brno Czech Republic

Molecular Biology Techniques Laboratory Faculty of Biology Adam Mickiewicz University in Poznan Umultowska 89 61 614 Poznań Poland

Muzeum Regionalne im Janusza Petera ul Zamojska 2 22 600 Tomaszów Lubelski Poland

National University of Kyiv Mohyla Academy Institute of Archaeology Hryhoriya Skovorody St 2 04655 Kyiv Ukraine

Polish Academy of Sciences Institute of Archaeology and Ethnology Sławkowska str 17 31 016 Kraków Poland

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